University of Groningen

Some aspects of the schooling behaviour of fish

Keenleyside, Miles Hugh Alston

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1955

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Keenleyside, M. H. A. (1955). Some aspects of the schooling behaviour of fish. [Thesis fully internal (DIV),
University of Groningen]. Martinus Nijhoff/Brill.

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ISIDE
;elf, has consummatorvvalue'
is thresholdlowering. Preven-
r an animal reacting to less and
ting in the so-called "vacuum
aen no external stimulus is aP-
r a c onsta n tstim ulus lT iNeen -
ce for the Presenceof internal
1 to respondto a given stimulus
r rudd (Table XVI) show that
oes not affect the intensitY of
getheragain. Holever, 'the need
t is clear.
nstinctive activities of the same
tivation of instinctive behaviour
a not h e rPa ttc r n" ( T r x r unc a N ,
ooling sticklebacksProvide con'
feeding motivation is correlated
' food, aithough theY grotlp more
iductive motivation leads to ces-
d femalesgroup less closelYthan
) closer schooling.The main dif-
he lack of suPPortfor assigning
rich can be comParedto the levels
ntres (seebelorn'-)'
he centralner\roussystem of fish
brain remo\:alhas lecl to cessation
6; WraeA'Lcr<,rg37) ancl has had
Flo'uvct'er,
1936;BenrvntN, rgttr ).
fortn it takcs arnong
arying in the
:r within one sPecies, is ind'irect
rechanismsin the central neÍ\rous
: socialinstinctin animals' "There
that are not Part of some instinct'
of a systernof centres controlling
when it strives to be in the neigh'
ies lvhen pcrformirlg somc' or all,
r, r95I, p. rrz). \'IovNrueN ancl
',Lonclntraprutctrt'lata)shor'vappe-
< when isolatedand also that these
6o 6r ASPECTS OF SCHOOLING BEHAVIOUR OF FISH 243
birds flock together when they are not engaged in any other activity. They
consider, therefore, that there is a. partially independent general social
motivation in some birds, comparableto incubation motivation, in which the
consummatory situation, as with flocking, consists of being in a certain
relation to objects in thc environrneut.
The question of rn'hetherthcre is a separatc schooling instinct can be
answered tentatively in the affirmative. Schooling behaviour of fish clearly
has some of the characteristicsof an instinctive activity, although cr.iclence
for the presenceof some other fcatures is inadequate.There is, holvever,
1ro superordinated,mystical "soci:rl centre" which is at a higher level than
other centres ancl rvhich controls a1i other types of behaviour. Schooling
should probably be placed at a lorv ler,,elin the organization of behaviour
because,while somc instincts (feeding, escapeand others) influence and
make use of schooling,the latter docs not directly influence other instincts.
SUtvf MARY
The purpose of this paper is to present a h1.-pothesison the nature of the schooling
behaviour of fish based on an ethological investigation of schooling. Recognizing the
disadvantages of a limited amount of data and of the use of different species for
different parts of the study, the following tentative picture is suggested.
Schooling may be considered an instinct as defined by TtwarncBrv and is at a rela-
tively low level in the hierarchical organization of behaviour. It has typical appetitive
behaviour and a consummatory situation. A single fish separated from its school searches
until perceiving a group of fish. It then approaches the group. In most cases vision is
the only sense involved in this approach. If more detailed specific stimuli are then
perceived (possibly through any of the sense organs) the fish ceases searching and
remains with the school; if not, it soon leaves, arrd appetitive behaviour continues until
the apploprizrte consullrnlatorv sitrration (being in il school of the same species) is
attained. This hypothesis is based on the following points:
r. A school of fish is an aggregation fomed when one fish reacts to others by
renraining near them.
z. Typical featurcs of Gasterostetts aculea,tus ancl Scartlíníu,s erytlropltthtthtnts
schools are: performance of the same activity at the same time by all fish, lack of
aggressivcness between members and equality of rank of all members.
3. Blinded S card,iniws fail to sl-row typical schooling but rernaiu in an area where
odours from other Scard,inius can be detected. This response may keep schools of this
species from scattering widely at night.
4. Visual perception of a school of fish releases approach in single Scard,ínius and
Pristella rid.dlei.
g. When presented with two clil-ferent-sized schools of their own species single
Gasterosteus, Scurdinius and Leu,cisctts nt,tiltts prefer the larger to the smaller group.
ó. A small Gasterosteu.s prefcrs six large to six small Gaslerosteus.
7. A single Gasterosteu.sprefers a school of its orvn species to a school oÍ Rhodeus
an'rart6 but shows no consistent preference u'hen either P5'gosteus ptmgititts or Leu-
ciscus are presented together with Gasterostetts.
8. A single Pristalla preiers a school of unoperated Pristella to a school rvith am-
putated dorsal fins.'Ihe dorsal fin with its conspicuous bl:rck patch is jerkecl nore
 244 M. H. A. KEENLEvsTDE 6z

rapidly after alarm. This structure ancl its special movement nrav be consiclerecl a
social releaser.
g. Increased feeding motivation leads to lirnited dispersal of a sclrool of Gas/cro.stcus.
The heacl-dorvn feeding posture is a signal attracting others in a school to a source of
food.
ro. Alarn causes an increase in density of a scirool of Gosterostcus.
rt. With increasing reproductive rnotivation rnale Gastcrostc2l-r cease schoolins arrcl
try to hold territories. ïlemales disperse to a limited extent.

ACKI{OWLEDGEME}íTS
This work was carried out during the trvo years the author was a guest at 'rhe
Zoological Laborator5', The University of Groningen. 1'he author rvishes to express his
sincere gratitude to Pro[essor G. P. Br\ERnNos, Director of the laboratorl', for n-uiking
it possible for him to study lhere and for guidance, advice anci criticisms rlurir-rg the
course of the work.
The ready cooperation and assistance of the members of the scientific and technical
staff of the laboratory are deeply appreciated.
Thanks are extended to Miss A. F. Tucxrn for reading anci criticizing parts of the
tnanuscript and to Dr H. F. R. PnecHrl for translating the sulnmary into Gernran.
Two Canadian Governmcnt Overseas Scholarships, awarclecl annually by the Ro1'al
Societl' of Canada, are gratefully acknou'ledged. \\iithout this financial assistance the
author r,ry'ould have been unable to study in The Netherlands.

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